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Bulletin of Insectology 63 (1): 137-152, 2010 ISSN 1721-8861

Bumble bees of the ( )

Aulo MANINO, Augusto PATETTA, Giulia BOGLIETTI, Marco PORPORATO Di.Va.P.R.A. - Entomologia e Zoologia applicate all’Ambiente “Carlo Vidano”, Università di Torino, ,

Abstract

A survey of bumble bees (Bombus Latreille) of the Susa Valley was conducted at 124 locations between 340 and 3,130 m a.s.l. representative of the whole territory, which lies within the Cottian Central , the Northern , and the South-eastern . Altogether 1,102 specimens were collected and determined (180 queens, 227 males, and 695 workers) belonging to 30 species - two of which are represented by two subspecies - which account for 70% of those known in Italy, demonstrating the particular value of the area examined with regard to environmental quality and biodiversity. Bombus soroeensis (F.), Bombus me- somelas Gerstaecker, Bombus ruderarius (Mueller), Bombus monticola Smith, Bombus pratorum (L.), Bombus lucorum (L.), (L.), and (L.) can be considered predominant, each one representing more than 5% of the collected specimens, 12 species are rather common (1-5% of specimens) and the remaining nine rare (less than 1%). A list of col- lected specimens with collection localities and dates is provided. To illustrate more clearly the altitudinal distribution of the dif- ferent species, the capture locations were grouped by altitude. 83.5% of the samples is also provided with data on the plant on which they were collected, comprising a total of 52 plant genera within 20 plant families. The eight predominant Bombus species point out remarkable differences in botanical choices, in fact only three genera (Epilobium, Onobrychis, and Trifolium) were vis- ited by all eight species, and showed very different frequencies between the species.

Key words: altitude, Bombus, flora visited, mountain species, .

Introduction these studies show however unequivocally that 5 of the 43 species recorded in Italy have not been found lately The genus Bombus sensu lato Latreille 1802 (Hymenop- anymore. tera Apidae), comprising about 240 species present in In Italy, although there are bumble bee subspecies most continents, is distributed mainly in areas of typical of the South and the islands, the number of taxa , Asia and North America with a cold and tem- present in the North is higher because of the species perate climate (Williams, 1998; Benton, 2006). The with a boreal-alpine distribution and of those living in bumble bees are the only social whose distribu- the mountains at medium and/or high altitudes, that can tion extends up to the Arctic regions, where thus be defined as “orophilous”. The distribution of are characterized by active nest temperature control and such species is clearly affected by the altitude, as al- increased pilosity that make them well adapted to the ready proved by Pittioni (1938) and confirmed by the cold (Pouvreau, 1984). very rich literature afterwards; a recent review of this As regards the altitudinal range, bumble bees are pre- matter is given by the papers by Iserbyt et al. (2008) and sent from sea level up to 4,250 m a.s.l., varying consid- Iserbyt (2009), and the references they quote. In Pied- erably in size and density of the pubescence. However, mont (North-western Italy), this phenomenon was high- although the species of high altitudes are generally more lighted for the "Waldensian Valleys" of the Cottian hairy, it is not clear if there is a correlation between Alps, on the basis of collections made between 1946 hairiness and altitude (Pouvreau, 1984). and 1970, by Comba (1960 and 1972) and Comba and Bumble bees are an important component of ecosys- Comba (2001). tems and their pollinating activity is essential for the The wide variability of geo-morphological and cli- conservation of many plant species. Really because their matic conditions of the Alps, and their level of anthro- presence depends on the abundance and diversity of pogenic disturbance, suggested extending the observa- flora, they are useful indicators of environmental health tions to a contiguous area. This was done by choosing (Macdonald, 2003). In the global decline in biodiversity, the Susa Valley for its extensive and complex hydro- bees are not the exception, unfortunately. The decline, graphic network, the significant elevation difference be- triggered by habitat fragmentation and loss of useful tween the valley and the highest peaks, the large exten- flora, is now documented in Europe especially for social sions of alpine meadows and the ease of access also to bee species (Williams, 1986; Westrich, 1989; Banaszak, high elevations. Moreover, the bumble bee fauna on the 1995; Biesmeijer et al., 2006; Fitzpatrick et al., 2007; morainic hills at the valley outlet in the high Piedmon- Kosior et al., 2007; Williams et al., 2007). tese plain had been recently investigated (Quaranta et Information available on the Italian bumble bee fauna al., 2004). is less than that relative to other countries; therefore it is The territory is located in the Central Cottian Alps, in difficult to point out changes in the distribution and in the Northern Cottian Alps, and the South-eastern Graian the abundance of bumble bees in Italy. Records up to Alps (Marazzi, 2005). On the basis of the geomor- 1995 were assembled by Intoppa et al. (1995) and were phological evolution, it can be divided into three seg- recently reviewed along with later research (Intoppa et ments: High Susa Valley upstream of , Middle al., 2009), but further investigations are much needed; Susa Valley between Oulx and Susa, and Lower Susa

Valley, between Susa and the valley outlet. In the main summer, consistent with the climatic conditions and the valley, which has a curved course with an approxi- growth of the flora, in 114 sites representative of the mately east-west direction, the river flows, whole territory (figure 1); 94 sites were visited only into which, leaving out the smaller hydrographic net- once, the others twice or more (table 1). On the occasion work, three secondary branches flow: the Dora di Bar- of each visit to each site, bumble bees showing different donecchia, fueled in its initial part by the streams com- colour patterns (Intoppa, 2000) were sampled; whenever ing from Vallée Etroite and Valle di Rochemolles; the possible some specimens of each colour pattern were Dora di Cesana, formed by the streams Ripa and collected. Thuras, and the stream . The river basin, cov- In the course of investigations conducted in Susa Val- ering an area of 1,261 km2, extends from 3,538 m a.s.l. ley over the years 1973, 1976-'77, 1984-'85 and 1993, of the - also placed in an eccentric posi- 89 specimens of bumble bees had been collected in 12 tion with respect to the course of the river - to 300 m sites. They were kept in the collection of Di.Va.P.R.A. a.s.l. of the valley outlet. The highest peaks, on which and the relative data were added and processed together the dorsal watershed separating the valley from the ad- with the 2001-2006 data to obtain a more complete spe- jacent basins is articulated, often include relief exceed- cies list for the valley. They are presented here even ing 3,000 m a.s.l.. though some were previously included in the catalogue by Intoppa et al. (1995): i.e. three records of 1973, relat- ing to the municipality of , and other specimens, Materials and methods collected in the years 1984-'85, reported with the ge- neric location “Valle di Susa”. The area in which the study was conducted includes the For each bumble bee, capture date and location were entire valley of the Dora Riparia, which is currently di- recorded; the latter was georeferenced to determine alti- vided into the mountain communities of Alta Valle di tude and UTM WGS84 co-ordinates. Where possible, Susa and Bassa Valle di Susa and Val Cenischia and the plant on which the bumble bee was foraging was politically in some territories belonging to (basin collected and determined by reference to Aeschimann et of , Col de Mongenèvre and Vallée Etroite). al. (2004). Some small areas, which are located along the water- The classification up to the generic level was made re- shed between the valley of the Dora Riparia and that of ferring to Michener (2000); for the subdivision into the , but included in the catchment area of the subgenera the simplified classification proposed by Wil- latter, have also been taken into account in the munici- liams et al. (2008) was followed, while for the identifi- pal districts of and . cation of species and subspecies and the relative no- Observations and collections were made from 2001 to menclature the work by Intoppa et al. (2009) was 2006 during visits that have taken place in spring and adopted.

Figure 1. Collection sites. Localities are numbered according to table 1; shaded areas are part of the French territory.

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Table 1. Collection sites, subdivided by municipal district (Italy) and mountain territory (France), with their altitude, UMT WGS 84 co-ordinates (East-North) 32N , and the indication of the year(s) they were visited (num- ber of visits in brackets).

Locality Site m a.s.l. UTM WGS 84 year (visits) ITALY 1984 (8) 1.1 Drubiaglio 340 375810 4994860 1985 (11) 2.1 Lago Patarè 2810 329800 4999300 2003 (1) 2.2 Lago Sommeiller 2980 330230 4999870 2003 (1) 2.3 Les Arnauds 1360 317970 4993050 2004 (2) 2.4 Melezet 1380 317630 4992400 1977 (1) 2.5 Monte Jafferau 2660-2800 324340 4994440 2004 (1) 2.6 Passo dei Fourneaux settentrionale 3130 331250 4998540 2006 (1) 2.7 Pian dei Frati 2580 328800 4998850 2003 (1) 2.8 Pian dei Morti 2380 327330 4999320 2003 (1) 2.9 Punta Colomion 2020 319750 4991300 2004 (1) 2.10 Rifugio Scarfiotti 2150 327120 5000050 2003 (1) 2.11 Rochemolles 1640-1760 323250 4998430 2004 (1) 2.12 Strada per il Sommeiller 2710 329320 4999070 2003 (1) 2.13 Vallon du Fond* 2730-2820 329770 4998280 2006 (1) 2.14 Vallon du Fond** 2970-3070 330850 4998460 2006 (1) 3.1 Borgone Borgone 400 361130 4998300 2001 (3) 4.1 Colle della Croce di Ferro 2500-2550 353390 5005410 2003 (1) 5.1 Caprie 380 368610 4997350 1985 (5) 5.2 Novaretto 360 370230 4996910 2002 (1) 6.1 Colle Bercia 2190 325070 4977050 2004 (1) 6.2 Fenils 1390 325780 4983450 2004 (1) 6.3 Grangia la Coche Alta 1930 324140 4978500 2004 (1) 6.4 Lago Fontana Fredda 2170 325150 4974140 2004 (1) 6.5 Lago Nero 2020 325940 4974480 2004 (1) 6.6 Monte Cruzeau 1570 326680 4978840 2004 (1) 6.7 Sagna Longa 2010-2080 325340 4977680 2004 (1) 7.1 Chianocco 540 355980 5000980 2002 (1) 7.2 Grangia Pianfè 1240 355580 5003530 2002 (1) 2001 (1) 7.3 Pavaglione 1010-1050 357370 5002310 2002 (1) 8.1 Cima Ciantiplagna 2650-2750 343140 4992850 2003 (1) 8.2 Frais 1460 342280 4996480 2004 (1) 9.1 Claviere 1750 322690 4978610 2004 (1) 10.1 Bertolere 940 365030 4999750 2002 (1) 10.2 Colle del Colombardo 1890 367310 5005180 2005 (1) 10.3 Colombardino 1890 366620 5005490 2005 (1) 10.4 Frassinere 980 364460 4999090 2003 (1) 1985 (2) 10.5 820 365530 4999570 2002 (1) 10.6 Mollette 1240 362560 5000040 2002 (2) 2002 (3) 10.7 Prarotto 1430 361440 5000980 2003 (1) 10.8 Punta Sbaron 2090 365050 5003950 2005 (1) 10.9 Tomba di Matolda 2010 365550 5004950 2005 (1) 11.1 Colle dell'Assietta 2450 339170 4992800 2004 (1) 11.2 Rifugio Levi-Molinari 1870 334440 4997900 2004 (1) 12.1 Colle Clapier 2480 336670 5003720 2005 (1) 12.2 Cresto 1070 342340 5000510 2003 (1) 12.3 Forte 690 344950 5000550 2002 (1) 12.4 Pra Piano 1500 341620 5001370 2003 (1) 12.5 Rio Clapier 2430 336280 5003420 2005 (1) 12.6 Sant'Andrea 810 343800 5000860 2001 (1) 2002 (1) 13.1 Pian Gelassa 1525-1540 345000 4995990 2003 (1) (Continued)

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(Table 1 Continued) Locality Site m a.s.l. UTM WGS 84 year (visits) 13.2 Punta del Mezzodì 2600-2650 344080 4993650 2003 (1) 14.1 Mattie Giordani 690 352250 4997930 2001 (1) 2002 (1) 15.1 2150-2250 346620 4992920 2003 (2) 2001 (1) 15.2 Corbolej 670 348460 4998130 2002 (3) 15.3 Piano del Tiraculo 1930 346950 4993710 2002 (1) 16.1 Alpe Arcella* 2030-2070 350830 5004700 2003 (1) 16.2 Alpe Arcella** 2110 351060 5004700 2003 (1) 2002 (1) 16.3 Il Trucco 1710-1740 347730 5003160 2003 (1) 16.4 La Riposa 2180 349290 5004530 2002 (2) 16.5 Monte Palon* 2200-2400 352110 5004320 2003 (1) 16.6 Monte Palon** 2460 352340 5004510 2003 (1) 1976 (1) 17.1 Moncenisio Ferrera Cenisio 1450 341510 5007640 2001 (1) 2001 (3) 18.1 Novalesa 910 343480 5006800 2002 (2) 18.2 San Rocco 740 344120 5004620 2002 (1) 19.1 Oulx Oulx 1060 330040 4989640 1993 (9) 19.2 C. Blanc 1130 324630 4990390 2004 (1) 19.3 C. Pourachet 2070 325450 4986220 2004 (1) 19.4 Chateau Beaulard 1410 324550 4988840 2004 (1) 19.5 Col Basset 2440-2510 327090 4993960 2004 (1) 19.6 Grotta dei Saraceni 2260 328370 4993840 2004 (1) 19.7 Monfol 1650 332610 4990190 1993 (1) 19.8 Monte Genevris 2480 334820 4988670 2004 (1) 19.9 Monte Vin Vert 2390 327910 4993550 2004 (1) 19.10 Rochers de l'Aigle 2620-2650 326390 4994400 2004 (1) 19.11 Savoulx 1130 325510 4990080 2004 (1) 19.12 Vazon 1680 327810 4987470 2004 (1) 20.1 Pragelato Lago dell'Assietta 2520 337930 4991830 2004 (1) 21.1 Rubiana Rubiana 680 373110 5000190 1973 (1) 21.2 Favella 990 371620 5001750 1973 (1) 22.1 Salbertrand 1040 334430 4993050 1993 (2) 22.2 Colle Blegier 2360 335830 4989190 2004 (1) 22.3 Colletto Pramand 2090 329900 4992780 2004 (1) 22.4 Forte Pramand 2140 330010 4992410 2004 (1) 23.1 San Giorio di Susa 440 357110 4998660 2001 (2) 1984 (1) 23.2 Malpasso 410 358960 4997930 1985 (7) 24.1 Sauze di Cesana 1550 330660 4978570 2004 (1) 24.2 Alpe Gran Mioul 2420 339720 4969510 2004 (1) 24.3 Argentera 1900 336950 4972680 2004 (1) 24.4 Brusà del Plan 1820 333950 4975400 2004 (1) 24.5 Grange Sises 1860 332050 4978960 2004 (1) 24.6 Rollières 1490 329310 4978240 2004 (1) 24.7 Valle del Gran Mioul 2110-2250 339160 4970520 2004 (1) 25.1 Sauze d'Oulx Colle di Costa Piana 2310 334540 4987440 2004 (1) 25.2 Lago Nero 2290 332100 4985220 2004 (1) 25.3 Notre Dame des Broussailles 2280-2320 331310 4985020 2004 (2) 25.4 Richardette 1710 332580 4988340 2004 (1) 26.1 Campo sportivo 2070 332620 4980860 2004 (1) 26.2 Plagnols 1970 332250 4979950 2004 (1) 27.1 Usseaux Colle della Vecchia 2420-2490 344630 4992980 2003 (1) 27.2 Colle delle Finestre 2000 346550 4991960 2003 (1) 27.3 Madonna del Dente 2460-2550 344570 4993190 2003 (1) 27.4 Monte Pintas 2330-2400 345530 4993130 2003 (2) (Continued)

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(Table 1 Continued) Locality Site m a.s.l. UTM WGS 84 year (visits) 28.1 Venaus 560 344850 5001190 2002 (2) 2002 (2) 28.2 Fondo di Bar 1590 340460 5006320 2003 (1) 28.3 Grangia della Vecchia 2200-2250 339970 5004370 2004 (1) 28.4 Grangia Marzo 1990-2080 341160 5004090 2004 (1) 28.5 Rio Tillirei 2000 340930 5003440 2004 (1) 28.6 Torrente Clanero 2120-2150 340410 5004360 2004 (1) 29.1 Villar Dora 350 372510 4996830 1977 (2) 29.2 Torre del Colle 350 372160 4995670 2001 (1) FRANCE 30.1 Mont Cenis Cumba di Crevacuore 2150 338000 5009110 2003 (1) 30.2 Barrage 2030 340150 5010220 2002 (1) 30.3 Fort Variselle 2040-2100 338410 5009590 2003 (1) 30.4 Gran Scala 1760 339590 5008880 2002 (2) 30.5 Grand-Croix 1850 339510 5009370 2002 (1) 30.6 Lac de Rotherel 1950 339690 5007460 2003 (1) 30.7 Mont Malamot* 2180-2400 337580 5008880 2003 (1) 30.8 Mont Malamot** 2500-2650 336620 5007900 2003 (1) 2001 (3) 30.9 Piano San Nicolao 1720-1770 339590 5008550 2002 (3) 2003 (1) 2001 (1) 30.10 Plan des Fontainettes 2090 338830 5012100 2002 (1) 31.1 Montgenèvre Col de Mongenèvre 1870 320760 4978130 2004 (1) 32.1 Valleé Etroite Grange di Valle Stretta 1760 312980 4993480 2004 (1) 32.2 La Grosse Somme 2150-2400 311810 4998170 2004 (1) 32.3 Vallon de 1820-2100 311870 4995650 2004 (1)

Results cumulative number of species identified vs the number of specimens (figure 3) tends to get asymptotic after the Altogether, 1,102 specimens (180 queens, 227 males, first 200 specimens, thus marking that the probability to and 695 workers) were collected in 124 sites between find further species in the Susa Valley is poor. 340 and 3,130 m a.s.l. (mean value 1.950 m). They be- To illustrate more clearly the altitudinal distribution of long to 30 species, eight of which can be considered pre- the different species, the collection sites were grouped dominant, each one representing over 5% of the speci- by altitude (table 2), following the indications by mens collected, 12 are common (1-5% of the specimens) Comba and Comba (2001). Neglecting the rare species, and the remaining nine are rare (less than 1%) (table 2). for which an evaluation is difficult to express, it is noted In the period 1973-1993, 15 species had already been that: (L.), Bombus rupestris (F.), identified, while the remaining 15 were added in the Bombus vestalis (Geoffroy), Bombus pratorum (L.), years 2001-2004 (figure 2) and no more species were Bombus lucorum (L.), Bombus terrestris (L.), and Bom- found in the last two years; the rarefaction curve of the bus lapidarius (L.) are characterized by a more or less

Figure 2. Number of Bombus specimens collected in the Susa Valley from 1973 to 2006 (dotted) and number of spe- cies identified for the first time in each year (black).

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142 Table 2. Number of Bombus specimens collected in the Susa Valley and number of queen (♀), male (♂), and workers ( ) sampled at different altitudes.

Specimens ≤700 701-1000 1001-1400 1401-1800 1801-2100 2101-2400 >2400 No. % ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀ ♂ B. mendax 35 3.18 6 2 3 11 13 B. soroeensis 83 7.53 1 1 1 11 10 1 2 22 3 23 8 B. subterraneus 5 0.45 2 1 1 1 B. argillaceus 1 0.09 1 B. hortorum 21 1.91 1 4 3 1 1 2 1 4 2 1 1 B. ruderatus 1 0.09 2 B. humilis 25 2.27 1 2 8 2 5 2 1 1 3 B. mesomelas 75 6.81 1 1 3 1 3 12 3 18 3 3 14 6 1 6 B. mucidus 13 1.18 1 3 2 1 4 2 B. pascuorum 36 3.27 6 11 4 2 2 1 1 9 B. ruderarius 73 6.62 3 10 5 14 11 14 4 3 8 1 B. sylvarum 18 1.63 4 14 B. bohemicus 18 1.63 1 7 3 1 4 1 1 B. flavidus 3 0.27 3 B. maxillosus 1 0.09 1 B. quadricolor 1 0.09 1 B. rupestris 18 1.63 4 1 5 1 1 1 2 1 2 B. sylvestris 7 0.64 1 2 1 1 1 1 B. vestalis 14 1.27 1 1 1 1 8 1 1 B. hypnorum 4 0.36 2 1 1 B. monticola 101 9.17 1 13 1 8 8 25 9 10 26 B. pratorum 56 5.08 2 5 2 2 1 11 1 16 11 5 B. pyrenaeus 46 4.17 2 1 1 8 3 1 18 1 6 5 B. alpinus 28 2.54 1 4 1 7 15 B. cryptaum 8 0.73 1 1 1 3 2 B. lucorum 170 15.42 2 1 2 1 3 2 4 9 17 1 8 19 9 33 2 14 43 B. terrestris 95 8.62 3 2 17 1 11 1 6 2 3 4 3 4 7 17 6 8 B. wurflenii 35 3.18 1 5 1 7 12 1 4 4 B. lapidarius 77 6.99 3 2 3 3 3 2 12 7 3 1 8 2 12 16 B. sichelii 34 3.09 2 2 2 7 2 2 11 3 3

mendax Gerstaecker and Bombus alpinus (L.), instead, are found, with only one exception, always above 2,100 m a.s.l. Altogether 83.5% of the samples have data on the plant on which they were collected; a total of 52 plant genera within 20 plant families were visited (figure 4). Among all the plants, the prevail (36.5% of the visits) with nine genera and some species visited with a high frequency: Onobrychis viciifolia Scop., Trifolium pra- tense L., Trifolium alpinum L., and Lotus alpinus (DC.) Schleicher. The Asteraceae represented 15.5% of the vis- its (82.5% were species of the genus Cirsium) and the Onagraceae represented 13.0% of the visits (only Epilo-

bium angustifolium L.). Less visited, but by no means Figure 3. Rarefaction curve, with 95% confidence in- the least, are the with 9.9% of the visits tervals, of the cumulative mumber of Bombus species (35.1% were Salvia pratensis L. and 42.9% were some identified in the Susa Valley. Computations and draw- species of the genus Thymus) and the Ericaceae with ing performed using PAST: Paleontological Statistics 5.2% of the visits thanks, almost exclusively (93.8%), to Software Package for Education and Data Analysis Rhododendron ferrugineum L.. The eight predominant (Hammer et al., 2001). Bombus species, referring to 631 observations out of the total 920 ones, visited plants belonging to 47 genera of all the 20 plant families recorded (table 3). But if we uniform presence throughout the area; Bombus soroeen- consider the plant genera visited by each of the eight sis (F.) and Bombus ruderarius (Muller) are absent be- Bombus species, we see that the range goes from 13 low 1,000 m a.s.l. and were preferably found at inter- plant genera visited by B. mesomelas to 28 plant genera mediate or medium-high elevations; Bombus monticola visited by B. terrestris and that there is a positive corre- Smith, Bombus pyrenaeus Perez, Bombus wurflenii Ra- lation between the number of plant visits of each Bom- doszkowski, and Bombus sichelii Radoszkowski do not bus species and the number of plant genera visited (Pear- occupy low elevations, but start to appear only above son Product Moment Correlation Coefficient = 0.733; P 1,400 m a.s.l., reaching the highest places; the same can = 0.0384). Nevertheless we point out some remarkable be said, with some exceptions, also for Bombus me- differences in botanic preferences of the different spe- somelas Gerstaecker and Bombus bohemicus Seidl; cies; only three genera (Epilobium, Onobrychis, and Tri- Bombus humilis Illiger, Bombus pascuorum (Scopoli), folium) were visited by all eight species and, however, and Bombus sylvarum (L.) do not exceed 1,400 m a.s.l. with very different frequencies. (the first two) and 1,800 m a.s.l. (the third one); Bombus

Figure 4. Number of plant genera (black) visited by bumble bees in the Susa Valley and number of Bombus specimens collected (dotted) from each plant family. Plant families are listed in a systematic order (Aeschimann et al., 2004).

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Table 3. Plant genera visited by the eight predominant Bombus species in the Susa Valley. Plant families are listed in a systematc order (Aeschimann et al., 2004) and diversity computations were performed using PAST: Paleon- tological Statistics Software Package for Education and Data Analysis (Hammer et al., 2001).

Plant family Plant genus B. soroeensis B. mesomelas B. ruderarius B. monticola B. pratorum B. lucorum B. terrestris B. lapidarius Ranunculaceae Ranunculus 1.2% Caryophyllaceae Silene 0.7% Polygonaceae Polygonum 1.3% 2.1% 2.4% Cistaceae Helianthemum 1.2% 3.5% 1.2% Ericaceae Rhododendron 6.4% 1.7% 9.7% 12.9% 3.5% 1.2% 1.9% Crassulaceae Sedum 1.3% 1.6% 2.4% 1.4% Saxifragaceae Saxifraga 2.1% Rosaceae Dryas 1.2% Geum 9.7% 9.2% Malus 1.2% 1.5% Potentilla 3.7% Prunus 1.4% 2.4% Rosa 1.9% Fabaceae Anthyllis 1.3% 3.4% 3.2% 1.4% 2.9% Astragalus 3.4% 1.6% 3.7% 1.4% 2.4% 1.5% Lotus 1.6% 22.1% 3.7% 4.2% 1.2% 10.1% Medicago 1.4% 1.2% Onobrychis 7.7% 12.1% 19.4% 20.8% 5.6% 2.1% 8.2% 11.5% Trifolium 5.1% 31.1% 19.4% 8.5% 5.6% 11.7% 34.3% 21.6% 1.2% 0.7% 1.2% 5.8% Onagraceae Epilobium 26.8% 17.3% 4.8% 4.9% 9.2% 20.7% 4.8% 4.4% Geraniaceae Geranium 0.7% Boraginaceae Cynoglossum 1.6% 9.2% Echium 2.6% 1.7% 1.6% 2.4% 3.7% 1.4% 4.8% Eritrichium 0.7% 1.5% Lamiaceae Ajuga 1.7% 1.2% 0.7% 2.4% 1.5% Prunella 6.6% Salvia 2.6% 5.2% 9.2% 4.2% 4.8% 5.8% Stachys 4.8% Thymus 8.5% 3.7% 9.0% 2.4% 1.2% 1.9% 4.2% Rhinanthus 1.3% 3.2% 3.7% 2.8% 2.4% 2.9% Veronica 1.2% Globulariaceae Globularia 1.2% Campanulaceae Campanula 9.0% 1.6% 2.4% 0.7% Phyteuma 6.4% 1.6% 1.9% 0.7% 1.2% 2.9% Caprifoliaceae Lonicera 1.3% 1.6% Dipsacaceae Knautia 2.6% 3.2% 2.4% 1.5% Scabiosa 1.2% 0.7% 1.2% Asteraceae Carlina 1.7% 1.3% 1.6% 1.2% 1.2% Cirsium 21.7% 17.3% 13.5% 1.9% 13.1% 5.9% 21.6% Echinops 1.7% Helianthus 1.7% 1.9% 1.2% Taraxacum 1.6% 2.4% 1.5% Liliaceae Asphodelus 1.3% 9.2% 2.8% 4.8% 1.5% number of plant genera 17 13 20 18 19 28 27 17 number of Bombus specimens 78 58 62 82 54 144 84 69 Shannon index 2.28 2.03 2.56 2.36 2.74 2.77 2.63 2.35 lower bootstrap (95% confidence) 1.93 1.62 2.11 2.01 2.37 2.48 2.15 1.99 upper bootstrap (95% confidence) 2.38 2.15 2.62 2.44 2.73 2.83 2.70 2.44

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List of collected specimens Bombus hortorum ssp. hortorum (L. 1761) AVIGLIANA: Drubiaglio, 24.VII.1984 1 , 27.VII.1984 1 , Genus Bombus Latreille 1802 31.VIII.1985 1 . BORGONE: 30.III.2001 1♀. CONDOVE: Frassinere, 20.V.2003 1 ; Mocchie, 9.VIII.1985 1 ; Subgenus MENDACIBOMBUS Skorikov 1914 Mollette, 31.V.2002 1♀ 1 . GIAGLIONE: Sant’Andrea, 20.V.2003 1 . GRAVERE: Pian Gelassa, 18.VI.2002 1♀, Bombus mendax ssp. mendax Gerstaecker 1869 19.VI.2003 4 . MEANA DI SUSA: Colle delle Finestre, BARDONECCHIA: Lago Pataré, 1.VIII.2003 1♀ 6♂♂ 4 ; 17.VII.2002 1♂. MONCENISIO: Ferrera Cenisio, Pian dei Frati, 1.VIII.2003 1♀ 4♂♂ 3 ; Pian dei Morti, 7.VIII.1976 1♂. MONT CENIS (France): Gran Scala, 1.VIII.2003 1♂; Strada per il Sommelier, 1.VIII.2003 1 ; 3.VII.2002 1♀. OULX: Col Basset, 23.VII.2004 1 . VE- Vallon du Fond*, 11.VII.2006 1♀ 1 . BUSSOLENO: Colle NAUS: Rio Tillirei, 27.VII.2004 2 . VILLAR DORA: della Croce di Ferro, 18.VII.2003 4 . MOMPANTERO: 20.IV.1977 1 . Monte Palona, 18.VII.2003 1♂. SALBERTRAND: Colle Blegier, 13.VII.2004 2♀♀. SAUZE DI CESANA: Valle del ssp. autumnalis (F. 1793) Gran Mioul, 15.VII.2004 1♀. USSEAUX: Madonna del AVIGLIANA: Drubiaglio, 11.VIII.1984 1 . Dente, 25.VII.2003 1♂. VALLÉE ETROITE (France): La Grosse Somme, 9.VII.2004 3♀♀. Subgenus THORACOBOMBUS Dalla Torre 1880

Subgenus KALLOBOMBUS Dalla Torre 1880 Bombus humilis ssp. quasimuscorum Vogt 1909 CESANA TORINESE: Monte Cruzeau, 29.VI.2004 1 . Bombus soroeensis ssp. proteus (Gerstaecker 1869) MEANA DI SUSA: Corbolej, 14.V.2002 1♀. NOVALESA: BARDONECCHIA: Lago Patarè, 1.VIII.2003 1 ; Lago 31.V.2002 1♀. OULX: 3.VI.1993 1♀; Chateau Beaulard, Sommeiller, 1.VIII.2003 1 ; Les Arnauds, 9.VII.2004 1 ; 29.VI.2004 1 . SALBERTRAND: 3.VI.1993 1♀. SAN GIORIO Rochemolles, 2.IX.2004 6♂♂ 1 . BUSSOLENO: Colle DI SUSA: Malpasso, 31.VII.1985 1 . SAUZE DI CESANA: della Croce di Ferro, 18.VII.2003 3 . CESANA TORI- 2.IX.2004 1 . VENAUS: Fondo di Bar, 18.VI.2002 1♀. NESE: Colle Bercia, 15.VII.2004 1 ; Fenils, 17.VI.2004 1♀; Sagna Longa, 15.VII.2004 2 . CHIOMONTE: Frais, Bombus humilis ssp. tristis Seidl 1837 27.VII.2004 1 . CLAVIERE: 2.IX.2004 2♂♂ 3 . AVIGLIANA: Drubiaglio, 27.VII.1984 1 , 12.VIII.1984 1 . CONDOVE: Colle del Colombardo, 30.VI.2005 4 ; Co- BARDONECCHIA: Les Arnauds, 9.VII.2004 1♂. CAPRIE: lombardino, 30.VI.2005 1♀ 1 ; Prarotto, 3.VII.2002 1 . 31.VII.1985 1 , 9.VIII.1985 2 , 28.VIII.1985 1♂, MEANA DI SUSA: Piano del Tiraculo, 17.VII.2002 1 . 30.VIII.1985 1♂. CHIANOCCO: Grangia Pianfè, 31.V.2002 MOMPANTERO: La Riposa, 22.VIII.2002 2♂♂; Monte Pa- 1♀; Pavaglione, 20.IX.2001 1♀. CONDOVE: Bertolere, lon*, 18.VII.2003, 1 . MONT CENIS (France): Fort 31.V.2002 1♀. GIAGLIONE: Cresto, 20.V.2003 1♀. OULX: Variselle, 8.VII.2003 1 ; Gran Scala, 3.VII.2002 1 , 20.VIII.1993 1♂. SALBERTRAND: 15.VII.1993 1 . SAN 22.VIII.2002 2 ; Piano San Nicolao, 31.V.2002 1♀. GIORIO DI SUSA: Malpasso, 9.VII.1985 1 , 9.VIII.1985 1 . MONTGENÈVRE (France): Col de Montgenèvre, 2.IX.2004 1 . OULX: C. Pourachet, 29.VI.2004 3 ; Col Basset, Bombus mesomelas ssp. mesomelas Gerstaecker 1869 23.VII.2004 1 ; Grotta dei Saraceni, 23.VII.2004 8 ; BARDONECCHIA: Lago Patarè, 1.VIII.2003 2♀♀ 1 ; Pian Roches de l’Aigle, 23.VII.2004 1 . PRAGELATO: Lago dei Frati, 1.VIII.2003 1♂ 1 ; Pian dei Morti, 1.VIII.2003 dell’Assietta, 13.VII.2004 1 . SALBERTRAND: Colle Ble- 1 ; Punta Colomion, 17.VI.2004 1 . CESANA TORINESE: gier, 13.VII.2004 1 ; Forte Pramand, 25.VI.2004 1 . Grangia la Coche Alta, 17.VI.2004 1 ; Lago Nero, SAUZE DI CESANA: 2.IX.2004 3♂♂ 1 . SAUZE D’OULX: 17.VI.2004 1♀. CHIOMONTE: Cima Ciantiplagna, Lago Nero, 13.VII.2004 3 ; Notre Dame des Brous- 25.VII.2003 1 . CONDOVE: Colombardino, 30.VI.2005 1 . sailles, 29.VI.2004 2 , 13.VII.2004 2 . SESTRIERE: MEANA DI SUSA: Colle delle Finestre, 17.VII.2002 2 , Plagnols, 2.IX.2004 4 . USSEAUX: Monte Pintas, 25.VII.2003 2 . MOMPANTERO: Alpe Arcella*, 25.VII.2003 2 . VENAUS: Grangia della Vecchia, 18.VII.2003 3 ; Alpe Arcella**, 18.VII.2003 1 ; La Ri- 27.VII.2004 1♂ 1 ; Grangia Marzo, 27.VII.2004 1♂; Rio posa, 17.VII.2002 2 , 22.VIII.2002 2♂♂; Monte Palon*, Tillirei, 27.VII.2004 1♂ 5 ; Torrente Clanero, 18.VII.2003 2 . MONCENISIO: Ferrera Cenisio, 27.VII.2004 1 . 15.VI.2001 1♀. MONT CENIS (France): Fort Variselle, 8.VII.2003 2 ; Gran Scala, 3.VII.2002 1♀, 22.VIII.2002 Subgenus SUBTERRANEOBOMBUS Vogt 1911 1♂; Grand-Croix, 18.VI.2002 3♀♀; Plan des Fontainettes, 22.VIII.2002 3♂♂ 7 . OULX: 20.VIII.1993 1 ; C. Bombus subterraneus ssp. latreillellus (Kirby 1802) Pourachet, 29.VI.2004 2♀♀; Monte Genevris, 13.VII.2004 AVIGLIANA: Drubiaglio, 12.VIII.1984 1 , 17.VIII.1984 1♀; Monte Vin Vert, 23.VII.2004 1♂. PRAGELATO: Lago 1 . MEANA DI SUSA: Colle delle Finestre, 17.VII.2002 dell’Assietta, 13.VII.2004 2♀♀. RUBIANA: 13.IX.1973 1 . 1♂. MONT CENIS (France): Grand-Croix, 18.VI.2002 1♀. SALBERTRAND: Colletto Pramand, 23.VII.2004 1 . SAUZE OULX: C.Blanc, 17.VI.2004 1♀. DI CESANA: 2.IX.2004 3 ; Alpe Gran Mioul, 15.VII.2004 1♀; Brusà del Pian, 25.VI.2004 1♀. SAUZE D’OULX: Notre Subgenus MEGABOMBUS Dalla Torre 1880 Dame des Broussailles, 29.VI.2004 2♀♀. SESTRIERE: Campo sportivo, 29.VI.2004 5♀♀. USSEAUX: Madonna del (Scopoli 1763) Dente, 25.VII.2003 3 ; Monte Pintas, 25.VI.2003 2 ; VILLAR DORA: 20.IV.1977 1♀. 25.VII.2003 2 . VALLÉE ETROITE (France): La Grosse

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Somme, 9.VII.2004 1♀; Vallon de Tavernette, 9.VII.2004 15.VII.1993 1 , 30.VII.1993 1 , 10.VIII.1993 2 , 2 . VENAUS: Fondo di Bar, 31.V.2002 1♀. 20.VIII.1993 5 , 28.VIII.1993 1 , 6.IX.1993 3 , 21.IX.1993 1 . Bombus mucidus ssp. mollis (Perez 1879) BARDONECCHIA: Pian dei Morti, 1.VIII.2003 1 ; Punta Subgenus Lepeletier 1832 Colomion, 17.VI.2004 1♀; Strada per il Sommeiller, 1.VIII.2003 1 . BUSSOLENO: Colle della Croce di Ferro, Bombus bohemicus ssp. bohemicus Seidl 1837 18.VII.2003 1♀. CESANA TORINESE: Lago Nero, BARDONECCHIA: Monte Jafferau, 23.VII.2004 1♀. CHIO- 17.VI.2004 2♀♀. CHIOMONTE: Cima Ciantiplagna, MONTE: Frais, 27.VII.2004 7♂♂. CONDOVE: Mollette, 25.VII.2003 1 . MONT CENIS (France): Piano San Nico- 3.VII.2002 1♂. MEANA DI SUSA: Colle delle Finestre, lao, 18.VI.2002 1♀. SAUZE DI CESANA: Alpe Gran Mioul, 25.VII.2003 1♀ 1♂. OULX: Grotta dei Saraceni, 15.VII.2004 1♀; Valle del Gran Mioul, 15.VII.2004 2♀♀. 23.VII.2004 3♂♂; Rochers de l’Aigle, 23.VII.2004 1♂. USSEAUX: Madonna del Dente, 25.VII.2003 2♀♀. SALBERTRAND: Colletto Pramand, 23.VII.2004 3♂♂.

Bombus pascuorum ssp. melleofacies Vogt 1909 Bombus flavidus ssp. alpium (Richards 1928) AVIGLIANA: Drubiaglio, 11.VIII.1984 1 , 12.VIII.1984 CHIOMONTE: Cima Ciantiplagna, 25.VII.2003 2♂♂. 1 , 24.VII.1985 1 , 30.VIII.1985 2 . BORGONE: MONT CENIS (France): Mont Malamot**, 8.VII.2003 1♂. 26.III.2001 1♀, 30.III.2001 2♀♀, 13.IV.2001 1♀.CAPRIE: 31.VII.1985 1 ; Novaretto, 19.IV.2002 1♀. CONDOVE: Bombus maxillosus ssp. maxillosus Klug 1817 Bertolere, 31.V.2002 1♀; Mocchie, 9.VIII.1985 1♀ 2 , VILLAR DORA: 20.IV.1977 1♀. 23.VIII.1985 1♀ 1♂; Mollette, 3.VII.2002 1 . GIAGLIONE: Cresto, 20.V.2003 1♀; Sant’Andrea, Bombus quadricolor ssp. meridionalis (Richards 1928) 20.V.2003 1♀ 1♂. GRAVERE: Pian Gelassa, 19.VI.2003 MOMPANTERO: Il Trucco, 17.VII.2002 1♀. 3 . MEANA DI SUSA: Corbolej, 13.IV.2001 1♀. MON- CENISIO: Ferrera Cenisio, 7.VIII.1976 3 . OULX: Cha- Bombus rupestris ssp. rupestris (F. 1793) teau Beaulard, 29.VI.2004 1 . SAN GIORIO DI SUSA: Mal- BARDONECCHIA: Pian dei Frati, 1.VIII.2003 1♂. CHIO- passo, 24.VII.1985 1 , 31.VII.1985 1 , 23.VIII.1985 1 , MONTE: Frais, 27.VII.2004 1♂. CONDOVE: Mollette, 29.VIII.1985 2 . SAUZE DI CESANA: 2.IX.2004 2 . 3.VII.2002 1♂. GIAGLIONE: Colle Clapier, 14.VII.2005 1♂. MEANA DI SUSA: Colle delle Finestre, 25.VII.2003 Bombus ruderarius ssp. ruderarius (Mueller 1776) 1♂. MONT CENIS (France): Gran Scala, 3.VII.2002 1♀; BARDONECCHIA: Les Arnauds, 17.VI.2004 1 ; Melezet, Piano San Nicolao, 29.VI.2001 2♀♀. OULX: Vazon, 31.VII.1977 1 ; Punta Colomion, 17.VI.2004 1 . CE- 29.VI.2004 1♀. SAN GIORIO DI SUSA: 15.V.2001 1♀, SANA TORINESE: Grangia la Coche Alta, 17.VI.2004 2♀♀; 25.V.2001 2♀♀. SAUZE DI CESANA: Valle del Gran Mi- Lago Nero, 17.VI.2004 1♀ 1 ; Sagna Longa, 15.VI.2004 oul, 15.VII.2004 1♀. VALLÉE ETROITE (France): La 3 . CHIANOCCO: Grangia Pianfé, 31.V.2002 1 . Grosse Somme, 9.VII.2004 1♀; Vallon de Tavernette, CONDOVE: Colle del Colombardo, 30.VI.2005 1 ; Co- 9.VII.2004 1♀. VENAUS: 14.V.2002 1♀; Fondo di Bar, lombardino, 30.VI.2005 2 . EXILLES: Rifugio Levi- 18.VI.2002 1♀; Rio Tillirei, 27.VII.2004 1♂. Molinari, 25.VI.2004 1 . GRAVERE: Pian Gelassa, 18.VI.2002 4♀♀ 2 , 19.VI.2003 2 . MEANA DI SUSA: Bombus sylvestris (Lepeletier 1832) Piano del Tiraculo, 17.VII.2002 1 . MOMPANTERO: Alpe CONDOVE: Prarotto, 31.V.2002 1♂. EXILLES: Rifugio Arcella**, 18.VII.2003 1 ; Il Trucco, 17.VII.2002 3♂♂ Levi-Molinari, 25.VI.2004 1♀. GIAGLIONE: Pra Piano, 2 , 20.V.2003 2♀♀; La Riposa, 17.VII.2002 6 , 20.V.2003 1♀. GRAVERE: Pian Gelassa, 18.VI.2002 1♀. 22.VIII.2002 3♂♂. MONCENISIO: Ferrera Cenisio, MEANA DI SUSA: Corbolej, 19.IV.2002 1♀. MONT CENIS 7.VIII.1976 1 , 15.VI.2001 1 . MONT CENIS (France): (France): Mont Malamot**, 8.VII.2003 1♂. USSEAUX: Gran Scala, 3.VII.2002 1 , 22.VIII.2002 1 ; Grand- Monte Pintas, 25.VII.2003 1♂. Croix, 18.VI.2002 1 ; Piano San Nicolao, 13.VII.2001 1 , 22.VIII.2002 1♂, 19.VI.2003 2 . OULX: C. Bombus vestalis ssp. obenbergeri (May 1944) Pourachet, 29.VI.2004 4♀♀; Col Basset, 23.VII.2004 1 ; CHIANOCCO: 19.IV.2002 1♀. CONDOVE: Mocchie, Vazon, 29.VI.2004 1 . SALBERTRAND: Colle Blegier, 31.V.2002 1♀; Mollette, 3.VII.2002 1♂; Prarotto, 13.VII.2004 1♀. SAUZE DI CESANA: Brusà del Plan, 3.VII.2002 1♂. OULX: Col Basset, 23.VII.2004 1♂. SAL- 25.VI.2004 1♀ 1 ; Rollières, 15.VII.2004 1♂; Valle del BERTRAND: Colletto Pramand, 23.VII.2004 2♂♂; Forte Gran Mioul, 15.VII.2004 1 . SAUZE D’OULX: Colle di Pramand, 25.VI.2004 1♀. VENAUS: Grangia Marzo, Costa Piana, 13.VII.2004 1♀; Notre Dames des Brous- 27.VII.2004 1♂; Rio Tillirei, 27.VII.2004 5♂♂. sailles, 13.VII.2004 1♀. SESTRIERE: Campo Sportivo, 29.VI.2004 2♀♀. VALLÉE ETROITE (France): Grange di Subgenus Dalla Torre 1880 Valle Stretta, 17.VI.2004 2♀♀; La Grosse Somme, 9.VII.2004 1♀; Vallon de Tavernette, 9.VII.2004 1♀ 2 . ssp. ericetorum (Panzer 1801) VENAUS: Fondo di Bar, 31.V.2002 2♀♀. CHIOMONTE: Frais, 27.VII.2004 2♂♂. OULX: C. Pourachet, 29.VI.2004 1 . Bombus sylvarum ssp. sylvarum (L. 1761) AVIGLIANA: Drubiaglio, 2.VIII.1984 1 , 24.VII.1985 1 , Bombus hypnorum ssp. hypnorum (L. 1758) 23.VIII.1985 1 . CAPRIE: 9.VIII.1985 1 . OULX: MEANA DI SUSA: Colle delle Finestre, 17.VII.2002 1 .

146

Bombus monticola ssp. hypsophilus (Skorikov 1912) Clapier, 14.VII.2005 1 . MONT CENIS (France): Fort BARDONECCHIA: Lago Patarè, 1.VIII.2003 1♂ 1 ; Monte Variselle, 8.VII.2003 2 ; Mont Malamot**, 8.VII.2003 Jafferau, 23.VII.2004 2 ; Pian dei Morti, 1.VIII.2003 1♂; Piano San Nicolao, 22.VIII.2002 1 . OULX: C. 1♂; Rifugio Scarfiotti, 1.VIII.2003 2♂♂; Strada per il Pourachet, 29.VI.2004 1 ; Grotta dei Saraceni, Sommeiller, 1.VIII.2003 2♀♀ 1 ; Vallon du Fond*, 23.VII.2004 1 . SALBERTRAND: Colle Blegier, 11.VII.2006 1 . BUSSOLENO: Colle della Croce di Ferro, 13.VII.2004 1♀ 1 . SAUZE DI CESANA: Argentera, 18.VII.2003 1♂ 2 . CHIOMONTE: Cima Ciantiplagna, 15.VII.2004 2 ; Valle del Gran Mioul, 15.VII.2004 1 . 25.VII.2003 1♀ 2 . CONDOVE: Tomba di Matolda, SAUZE D’OULX: Lago Nero, 13.VII.2004 1 ; Notre Dame 30.VI.2005 1 . GIAGLIONE: Rio Clapier, 14.VII.2005 des Broussailles, 13.VII.2004 2 . USSEAUX: Madonna 11 . GRAVERE: Punta del Mezzodì, 25.VII.2003 3♂♂ del Dente, 25.VII.2003 2♂♂ 1 . VALLÉE ETROITE 1 . MEANA DI SUSA: Colle delle Finestre, 17.VII.2002 (France): La Grosse Somme, 9.VII.2004 2♀♀ 6 . VE- 9 , 25.VII.2003 1♂ 6 ; Piano del Tiraculo, NAUS: Grangia della Vecchia, 27.VII.2004 2 ; Rio Til- 17.VII.2002 1 . MOMPANTERO: La Riposa, 17.VII.2002 lirei, 27.VII.2004 1 . 4 , 22.VIII.2002 3♂♂; Monte Palon*, 18.VII.2003 1 . MONCENISIO: Ferrera Cenisio, 15.VI.2001 1 . MONT CE- Subgenus ALPINOBOMBUS Skorikov 1914 NIS (France): Barrage, 3.VII.2002 1 ; Fort Variselle, 8.VII.2003 4 ; Gran Scala, 3.VII.2002 3 , Bombus alpinus ssp. helleri Dalla Torre 1882 22.VIII.2002 1♂; Lac de Rotherel, 19.VI.2003 1 ; Mont BARDONECCHIA: Lago Patarè, 1.VIII.2003 1 ; Lago Malamot*, 8.VII.2003 2 ; Mont Malamot**, 8.VII.2003 Sommeiller, 1.VIII.2003 5♂♂; Pian dei Frati, 1.VIII.2003 2♂♂ 1 ; Piano San Nicolao, 18.VI.2002 1 , 19.VI.2003 1 ; Strada per il Sommeiller, 1.VIII.2003 1♀; Vallon du 4 , Plan des Fontainettes, 22.VIII.2002 1♂. USSEAUX: Fond*, 11.VII.2006 2 ; Vallon du Fond**, 11.VII.2006 Colle della Vecchia, 25.VI.2003 1♂ 1 ; Madonna del 1♂ 1. BUSSOLENO: Colle della Croce di Ferro, Dente, 25.VI.2003 2 , 25.VII.2003 6♀♀ 2♂♂ 1 ; 18.VII.2003 1 . GIAGLIONE: Colle Clapier, 14.VII.2005 Monte Pintas, 25.VI.2003 2 ; 25.VII.2003 1♂ 1 . VE- 5 ; Rio Clapier, 14.VII.2005 4 ; MONT CENIS NAUS: Fondo di Bar, 18.VI.2002 2 , 20.V.2003 2 . (France): Fort Variselle, 8.VII.2003 1 ; Mont Malamot*, 8.VII.2003 3 . OULX: Rochers de l'Aigle, 23.VII.2004 Bombus pratorum ssp. pratorum (L. 1761) 1♂. VALLÉE ETROITE (France): La Grosse Somme, BARDONECCHIA: Les Arnauds, 9.VII.2004 1 ; Monte Jaf- 9.VII.2004 1 . ferau, 23.VII.2004 1 . CESANA TORINESE: Sagna Longa, 15.VII.2004 1 . CHIOMONTE: Frais, 27.VII.2004 1♂. Subgenus BOMBUS Latreille 1802 CLAVIERE: 2.IX.2004 1 . CONDOVE: Colle del Co- lombardo, 30.VI.2005 3 ; Colombardino, 30.VI.2005 Bombus cryptarum ssp. reinigianus Rasmont 1984 1 ; Frassinere, 20.V.2003 3 ; Mocchie, 31.V.2002 1 ; CESANA TORINESE: Lago Fontana Fredda, 15.VII.2004 Mollette, 31.V.2002 1 ; Prarotto, 20.V.2003 1 ; Punta 1 . CLAVIERE: 2.IX.2004 1♂. OULX: Monte Genevris, Sbaron, 30.VI.2005 1 ; Tomba di Matolda, 30.VI.2005 13.VII.2004 1 . PRAGELATO: Lago dell’Assietta, 1 . EXILLES: Rifugio Levi-Molinari, 25.VI.2004 1 . 13.VII.2004 1 . SALBERTRAND: Colle Blegier, GIAGLIONE: Rio Clapier, 14.VII.2005 2 . GRAVERE: 13.VII.2004 1 . SESTRIERE: Plagnols, 2.IX.2004 1♂. Pian Gelassa, 18.VI.2002 2♀♀ 3 . MEANA DI SUSA: VALLÉE ETROITE (France): La Grosse Somme, 9.VII.2004 Corbolej, 31.V.2002 1 . MONT CENIS (France): Gran 1 ; Vallon de Tavernette, 9.VII.2004 1 . Scala, 3.VII.2002 2 , 22.VIII.2002 1 ; Piano San Nico- lao,19.VI.2003 1 . OULX: C. Pourachet, 29.VI.2004 1♂ Bombus lucorum ssp. lucorum (L. 1761) 6 ; Col Basset, 23.VII.2004 1 ; Vazon, 29.VI.2004 1 . BARDONECCHIA: Lago Patarè, 1.VIII.2003 1♂ 3 ; Lago RUBIANA: Favella, 13.IX.1973 1 . SALBERTRAND: Forte Sommeiller, 1.VIII.2003 1 ; Les Arnauds, 9.VII.2004 Pramand, 25.VI.2004 5 . SAUZE DI CESANA: 2.IX.2004 3♂♂; Monte Jafferau, 23.VII.2004 1♂ 3 ; Pian dei 1 . SAUZE D’OULX: Notre Dame des Broussailles, Frati, 1.VIII.2003 5 ; Punta Colomion, 17.VI.2004 1 ; 29.VI.2004 2 , 13.VII.2004 1 . USSEAUX: Madonna del Rochemolles, 2.IX.2004 1♂ 1 ; Strada per il Sommeiller, Dente, 25.VII.2003 1 . VALLÉE ETROITE (France): La 1.VIII.2003 7 ; Vallon du Fond*, 11.VII.2006 3 ; Grosse Somme, 9.VII.2004 2 ; Vallon de Tavernette, Vallon du Fond**, 11.VII.2006 5 . BUSSOLENO: Colle 9.VII.2004 1 . VENAUS: 31.V.2002 1 ; Grangia della della Croce di Ferro, 18.VII.2003 3♂♂ 2 . CESANA Vecchia, 27.VII.2004 1 ; Grangia Marzo, 27.VII.2004 1 . TORINESE: Lago Fontana Fredda, 15.VII.2004 2 . CHIOMONTE: Cima Ciantiplagna, 25.VII.2003 2♂♂; Frais, Bombus pyrenaeus ssp. tenuifasciatus Vogt 1909 27.VII.2004 1♀1♂ 1 . CLAVIERE: 2.IX.2004 1♂ 1 . BARDONECCHIA: Lago Patarè, 1.VIII.2003 1 ; Lago CONDOVE: Colombardino, 30.VI.2005 2 ; Mollette, Sommeiller, 1.VIII.2003 1♂; Passo dei Fourneaux setten- 31.V.2002 1 ; Prarotto, 18.VI.2002 1♀, 3.VII.2002 1 ; trionale, 11.VII.2006 1♀; Pian dei Frati, 1.VIII.2003 2♂♂; Punta Sbaron, 30.VI.2005 1 . EXILLES: Colle Pian dei Morti, 1.VIII.2003 1♂ 1 ; Punta Colomion, dell’Assietta, 13.VII.2004 1 ; Rifugio Levi-Molinari, 17.VI.2004 1 ; Rifugio Scarfiotti, 1.VIII.2003 3 . BUS- 25.VI.2004 1 . GIAGLIONE: Colle Clapier, 14 VII 2005 SOLENO: Colle della Croce di Ferro, 18.VII.2003 1 . CE- 1 Rio Clapier, 14.VII.2005 6 . GRAVERE: Pian Ge- SANA TORINESE: Sagna Longa, 15.VII.2004 1 . CHIO- lassa, 19.VI.2003 1 ; Punta del Mezzodì, 25.VII.2003 1♂ MONTE: Cima Ciantiplagna, 25.VII.2003 1 ; Frais, 1 . MEANA DI SUSA: Colle delle Finestre, 17.VII.2002 27.VII.2004 1♂. CLAVIERE: 2.IX.2004 1♂. EXILLES: Ri- 5 , 25.VI.2003 4 , 25.VII.2003 2♂♂ 2 ; Corbolej, fugio Levi-Molinari, 25.VI.2004 1♂. GIAGLIONE: Colle 31.V.2002 1 ; Piano del Tiraculo, 17.VII.2002 1♂. MON-

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PANTERO: Il Trucco, 17.VII.2002 2 ; La Riposa, Somme, 9.VII.2004 1 ; Vallon de Tavernette, 9.VII.2004 22.VIII.2002 1♂; Monte Palon*, 18.VII.2003 3 . MON- 1 . VENAUS: 31.V.2002 2 ; Rio Tillirei, 27.VII.2004 CENISIO: Ferrera Cenisio, 15.VI.2001 1 . MONT CENIS 1♂. VILLAR DORA: 26.III.1977 1♀, 20 IV 1977 1 . (France): Fort Variselle, 8.VII.2003 1 ; Gran Scala, 3.VII.2002 1♀ 1 , 22.VIII.2002 1 ; Mont Malamot*, Subgenus ALPIGENOBOMBUS Skorikov 1914 8.VII.2003 5 ; Piano San Nicolao, 29.VI.2001 1♀, 22.VIII.2002 1 , 19.VI.2003 1 ; Plan des Fontainettes, Bombus wurflenii ssp. mastrucatus (Gerstaecker 1869) 28.VIII.2001 1♀, 22.VIII.2002 3♂♂ 5 . MONT- BARDONECCHIA: Lago Patarè, 1.VIII.2003 1 ; Pian dei GENÈVRE (France): Col de Montgenèvre, 2.IX.2004 3 . Frati, 1.VIII.2003 2 ; Pian dei Morti, 1.VIII.2003 1♂ 1 ; NOVALESA: 20.IX.2001 1♀. OULX: Col Basset, Punta Colomion, 17.VI.2004 1 ; Strada per il Sommeiller, 23.VII.2004 2♂♂ 2 ; Monte Genevris, 13.VII.2004 1 ; 1.VIII.2003 1 . MOMPANTERO: La Riposa, 22.VIII.2002 Salvoulx, 2.IX.2004 1 ; Vazon, 29.VI.2004 1 . PRAGE- 1 . MONCENISIO: Ferrera Cenisio, 7.VIII.1976 1♂. MONT LATO: Lago dell’Assietta, 13.VII.2004 2 . SALBER- CENIS (France): Barrage, 3.VII.2002 1♀ 1 ; Cumba di TRAND: Colle Blegier, 13.VII.2004 2♂♂ 3 ; Forte Pra- Crevacuore, 8.VII.2003 1 ; Fort Variselle, 8.VII.2003 mand, 25.VI.2004 3 . SAN GIORIO DI SUSA: Malpasso, 2 ; Lac de Rotherel, 19.VI.2003 1 ; Mont Malamot*, 24.VII.1984 1 , 15.VIII.1985 1♂. SAUZE DI CESANA: 8.VII.2003 1 ; Piano San Nicolao, 19.VI.2003 4 ; Plan 2.IX.2004 6♂♂ 3 ; Grange Sises, 2.IX.2004 1 . SAUZE des Fontainettes, 22.VIII.2002 7♂♂ 7 . SAUZE DI CE- D’OULX: Colle di Costa Piana, 13.VII.2004 1 . SESTRI- SANA: 2.IX.2004 1 . ERE: Plagnols, 2.IX.2004 1♂ 2 . USSEAUX: Colle delle Finestre, 25.VII.2003 1♂ 1; Madonna del Dente, Subgenus MELANOBOMBUS Dalla Torre 1880 25.VII.2003 2♀♀ 4♂♂; Monte Pintas, 25.VI.2003 4 , 25.VII.2003 1 . VALLÉE ETROITE (France): Grange di Bombus lapidarius ssp. lapidarius (L. 1758) Valle Stretta, 17.VI.2004 1 ; Vallon de Tavernette, AVIGLIANA: Drubiaglio, 23.VII.1985 1 . BARDONEC- 9.VII.2004 1 . VENAUS: Grangia della Vecchia, CHIA: Les Arnauds, 17.VI.2004 1♀, 9.VII.2004 1 ; Passo 27.VII.2004 3♂♂; Rio Tillirei, 27.VII.2004 2♂♂; Tor- dei Fourneaux settentrionale, 11.VII.2006 1 ; Pian dei rente Clanero, 27.VII.2004 1♂. VILLAR DORA:Torre del Frati, 1.VIII.2003 2 ; Strada per il Sommeiller, Colle, 26.III.2001 2♀♀. 1.VIII.2003 1 . BUSSOLENO: Colle della Croce di Ferro, 18.VII.2003 1 . CAPRIE: Novaretto, 19.IV.2002 1 . Bombus terrestris ssp. terrestris (L. 1758) CHIANOCCO: Grangia Pianfé, 31.V.2002 2♀♀ 1 . CHIO- AVIGLIANA: Drubiaglio, 27.VII.1984 1♂, 12.VIII.1984 MONTE: Cima Ciantiplagna, 25.VII.2003 3 ; Frais, 1 , 28.VI.1985 1 , 12 VII 1985 1 , 4.VIII.1985 1 , 27.VII.2004 3♂♂. CLAVIERE: 2.IX.2004 1♂. CONDOVE: 7.VIII.1985 1 , 9.VIII.1985 1 , 31.VIII.1985 1♂ 1 . Bertolere, 31.V.2002 1 ; Colle del Colombardo, BARDONECCHIA: Lago Patarè, 1.VIII.2003 1 ; Punta Co- 30.VI.2005 3 ; Prarotto, 31.V.2002 2♀♀, 18.VI.2002 lomion, 17.VI.2004 1 ; Strada per il Sommeiller, 4♀♀ 1, 3.VII.2002 1♀ 1♂; Tomba di Matolda, 1.VIII.2003 1 . BUSSOLENO: Colle della Croce di Ferro, 30.VI.2005 1 . GIAGLIONE: Colle Clapier, 14.VII.2005 18.VII.2003 1♂ 1 . CAPRIE: 28.VI.1985 1 , 30.VIII.1985 2 ; Sant’Andrea, 20.V.2003 1 . GRAVERE: Punta del 1 ; Novaretto, 19.IV.2002 2♀♀. CHIANOCCO: 19.IV.2002 Mezzodì, 25.VII.2003 1 . MATTIE: Giordani, 25.V.2001 1 ; Grangia Pianfè, 31.V.2002 3 ; Pavaglione, 1♀. MEANA DI SUSA: Colle delle Finestre, 25.VI.2003 1 , 3.VII.2002 4 . CHIOMONTE: Cima Ciantiplagna, 25.VII.2003 2 . MONPANTERO: Alpe Arcella*, 25.VII.2003 2♂♂ 1 . CONDOVE: Bertolere, 31.V.2002 18.VII.2003 2 ; Il Trucco, 20.V.2003 1 ; La Riposa, 2 ; Frassinere, 20.V.2003 1 ; Mollette, 31.V.2002 1 ; 17.VII.2002 2♀♀ 3 ; Monte Palon**, 18.VII.2003 1 . Prarotto, 18.VI.2002 2♂♂ 1. GIAGLIONE: Cresto, MONT CENIS (France): Gran Scala, 22.VIII.2002 1♂; 20.V.2003 1 ; Forte, 14.V.2002 2 ; Pra Piano, Grand-Croix, 18.VI.2002 1♀; Mont Malamot*, 20.V.2003 1♀. MEANA DI SUSA: Colle delle Finestre, 8.VII.2003 1 ; Piano San Nicolao, 15.VI.2001 1♀; Plan 17.VII.2002 1♂ 1 , 25.VII.2003 2♂♂ 3 ; Corbolej, des Fontainettes, 22.VIII.2002 1 . NOVALESA: 14.V.2002 1 . MOMPANTERO: Alpe Arcella*, 15.VI.2001 1♀ 1 , 29.VI.2001 1♀, 18.VI.2002 1♀. 18.VII.2003 2 ; La Riposa, 17.VII.2002 3♂♂ 3 ; OULX: Grotta dei Saraceni, 23.VII.2004 2 ; Vazon, 29 Monte Palon*, 18.VII.2003 1♂ 2 ; Monte Palon**, VI 2004 1♀. PRAGELATO: Lago dell’Assietta, 13.VII.2004 18.VII.2003 1♂. MONCENISIO: Ferrera Cenisio, 1 . SALBERTRAND: Forte Pramand, 25.VI.2004 1 . 15.VI.2001 2 . MONT CENIS (France): Gran Scala, SAUZE DI CESANA: 2.IX.2004 1♂. SAUZE D’OULX: Rich- 22.VIII.2002 1♂; Mont Malamot*, 8.VII.2003 1 ; Piano ardette, 13.VII.2004 1 . USSEAUX: Madonna del Dente, San Nicolao, 19.VI.2003 1 . NOVALESA: 15 VI 2001 1 , 25.VI.2003 1 , 25.VII.2003 2 ; Monte Pintas, San Rocco, 14.V.2002 3 . OULX: 22.IV.1993 1♀; Mon- 25.VI.2003 1 , 25.VII.2003 1 . VENAUS: 31.V.2002 1♀; fol, 22.IV.1993 1♀; Savoulx, 2.IX.2004 1 . PRAGELATO: Fondo di Bar, 31.V.2002 2♀♀, 18.VI.2002 1♀; Rio Til- Lago dell’Assietta, 13.VII.2004 2 . RUBIANA: Favella, lirei, 27.VII.2004 1 . VILLAR DORA: 20.IV.1977 1♀. 13.VII.1973 1♂. SALBERTRAND: Forte Pramand, 25.VI.2004 4 . SAN GIORIO DI SUSA: Malpasso, Bombus sichelii ssp. flavissimus Tkalcu 1974 9.VIII.1985 1 . SAUZE D’OULX: Colle di Costa Piana, BARDONECCHIA: Lago Patarè, 1.VIII.2003 2 ; Pian dei 13.VII.2004 1 . USSEAUX: Colle della Vecchia 25 VI Frati, 1.VIII.2003 3♂♂ 1 ; Pian dei Morti, 1.VIII.2003 2003 1 ; Colle delle Finestre, 25.VII.2003 2♂♂; 1♂ 1 ; Rifugio Scarfiotti, 1.VIII.2003 5 . CLAVIERE: Madonna del Dente, 25.VII.2003 2♂♂ 1 ; Monte Pintas, 2.IX.2004 1 . MEANA DI SUSA: Colle delle Finestre, 25.VI.2003 1 . VALLÉE ETROITE (France): La Grosse 17.VII.2002 1 . MOMPANTERO: La Riposa, 22.VIII.2002

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1♂. MONT CENIS (France): Fort Variselle, 8.VII.2003 was found neither in the Susa Valley nor in the contigu- 2 ; Gran Scala, 3.VII.2002 1 , 22.VIII.2002 2♂♂; Plan ous “Waldensian Valleys” (Comba, 1960; 1972; Comba des Fontainettes, 22.VIII.2002 2♂♂ 4 . MONTGENÈVRE and Comba, 2001), while it is widespread in the French (France): Col de Montgenèvre, 2.IX.2004 1 . SAUZE DI Alps (Rasmont, 1988), where it was collected also in CESANA: Valle del Gran Mioul, 15.VII.2004 1♀. US- localities a few kilometers far from the Susa Valley SEAUX: Monte Pintas, 25.VI.2003 1 , 25.VII.2003 1 . (Delmas, 1962; Tkalců, 1973). On the other hand, the VALLÉE ETROITE (France): La Grosse Somme, 9.VII.2004 only record of this species in is doubtful 1♀ 1 . VENAUS: Grangia della Vecchia, 27.VII.2004 1 . (Amiet, 1996) and the first and only Italian records (In- toppa et al., 1995) are the much southern ones by Tkalců (1973) concerning two cols between Italy and Discussion and conclusions France, the Col de Tende and the Colle della Madda- lena; one must not forget also Bombus pomorum (Pan- The bumble bee species in the Susa Valley account for zer) and Bombus magnus Vogt, that are present in the 70% of those known in Italy, demonstrating the value of (Rasmont, 1988), but not in Switzerland the examined area with regard to environmental quality (Amiet, 1996) neither, presently as far as we know, in and biodiversity. This is very important and significant Italy (Intoppa et al., 2009). because of the primary role played by bumble bees in The distribution in Italy of the bumble bee subspecies maintaining the ecosystem through their pollinating ac- is still uncertain due also to the presence of individuals tivity. It is therefore disconcerting that so far the infor- with intermediate characters (Intoppa et al., 2009); the mation on the bumble bee fauna of the valley was scarce presence of B. pascuorum melleofacies in the Susa Val- and fragmentary (table 4), even taking into account the ley - a subspecies known until now only in the Italian reports by Comba (1972) on the side of Colle delle Fi- peninsula (Rasmont, 1983) - is noteworthy, but also nestre facing the Chisone Valley. Bombus pascuorum floralis (Gmelin) has been recorded All the species collected in the Susa Valley, except B. in only lately (Barbattini et al., 2007). maxillosus, were already known from (In- The scanty data on Italian bumble bee fauna suggest toppa et al., 1995), but 16 are reported now for the first the outmost caution about actual species and subspecies time for the valley. On the contrary, Bombus inexspec- distributions and should also foster further investiga- tatus (Tkalcu), Bombus barbutellus (Kirby), and Bom- tions in the large areas for which information is poor or bus jonellus (Kirby) were not found during the present completely lacking. Therefore it seemed preferable not investigation, while they had been recorded by previous to extend the comparison to other mountain ranges and researchers (table 4); the first species, extremely rare, neither to the outer side of the Alps, but to limit it to seems to have a parasitic behaviour upon B. ruderarius what is known on the Italian inner side, referring to the (Müller, 2006), the second one parasitizes the nests of catalogue by Intoppa et al. (1995) and the subsequent B. hortorum, not particularly frequent in the Susa Val- studies (Intoppa et al., 1999; Comba and Comba, 2001; ley, while the third one, mainly linked to Ericaceae, is 2005; Barbattini et al., 2007; Manino et al., 2007; Cor- rare in Italy (Intoppa et al., 1995; Barbattini, 2007) even nalba, 2009). Altogether, the following differences can though it could be more common than it seems (Intoppa be pointed out: a lack of recent records for B. alpinus et al., 1999). outside the Piedmont region; absence of Bombus It is most remarkable that Bombus brodmannicus Vogt mucidus Gerstaecker in and Venetia, while it

Table 4. species recorded in localities of the Susa Valley by previous researchers.

Gribodo, 1873 Comba, 1972 Pagliano, 1994 B. mendax Colle del Moncenisio Colle delle Finestre Sauze d’Oulx B. soroeensis Colle delle Finestre Colle delle Finestre B. humilis Monte Musinè B. inexspectatus Colle delle Finestre B. mesomelas Sauze d’Oulx Colle delle Finestre B. ruderarius Colle delle Finestre Pietrabianca di Bussoleno B. barbutellus Colle delle Finestre B. bohemicus Pietrabianca di Bussoleno B. flavidus Sauze d’Oulx B. quadricolor Sauze d’Oulx B. rupestris Colle delle Finestre B. sylvestris Colle delle Finestre B. vestalis Sauze d’Oulx B. jonellus Susa surroundings B. monticola Sauze d’Oulx B. pratorum Sauze d’Oulx B. lucorum Sauze d’Oulx

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is rare in Friuli-Venezia Giulia; the absence of B. men- families already reported by other authors (i.e. Alford, dax and B. sichelii and a relative rarity of B. pyrenaeus 1975; Rasmont, 1988; Benton, 2006; Iserbyt et al., in Venetia and Friuli-Venezia Giulia, while they are 2008). The poor number of observations and the lack of rather common both in Piedmont and Lombardy. On the data regarding the relative abundance/area covered by contrary, in addition to the above mentioned B. horto- the flora where and when the bumble bees were col- rum, also Bombus hypnorum (L.), B. humilis, and, at lected do not permit to draw meaningful conclusions least for Friuli-Venezia Giulia, B. sylvarum seem less about the botanical preferences of the rare species or common in the Susa Valley than elsewhere. The pres- even of the common ones; for the eight predominant ence of the subgenus Psithyrus is instead everywhere species it seems instead evident that, though they visit quite low and obviously in relation to that of the bumble mostly the more abundant plant genera in their foraging bee species they are nest-parasites of. areas, they show remarkably different preferences, even The altitudinal distribution of the species, highlighted if not always in line with what was pointed out by other in the Susa Valley, is broadly in agreement with what authors. In particular, we must underline that, as it was was recently detected in Lombardy by Intoppa et al. already pointed out by Rasmont (1988), B. soroeensis (1999) in the Adamello Natural Park, located in the visits the Campanulaceae much more assiduously than Adamello-Presanella-Alps (Marazzi, 2005), and by the other species. Cornalba (2009) in the upper Val Brembana, located in Without sufficiently comprehensive data on the past the Orobie Alps (Marazzi, 2005). Of the 21 species presence of bumble bees in the Susa Valley, it is impos- found in the three territories, which have rather different sible to establish if there has been a decrease in the geomorphological and vegetational characteristics, only number of species present in the valley and/or a shrink- B. soroeensis and B. lapidarius reach, and in a not in- ing in number or area, but hints of the occurrence of this significant number, remarkably higher elevations in the phenomenon do exist. The global climatic change that is Susa Valley. When comparing, however, the distribu- taking place and the consequent vegetation changes tion of species and their abundance at various altitudinal pose however a serious threat also to bumble bee com- belts as emerged from this investigation with what was munities; in particular a further increase in temperature observed between 1946 and 1970 in the “Waldensian would mean that the distribution area of the typical high Valleys” (Comba and Comba, 2001), it is clear that in mountain species would go through a drastic reduction the Susa Valley about half of the 25 species that are and fragmentation that could seriously impair their sur- common to both areas, are present in great numbers at a vival. significantly higher elevation; besides for B. soroeensis and B. lapidarius, which have already been mentioned earlier, and B. mendax, B. monticola, and B. alpinus, Acknowledgements which were already dealt with elsewhere (Manino et al., 2007), the phenomenon is particularly evident for B. We thank Maria Gioia Piazza and Francesco Intoppa for mesomelas, B. ruderarius, and B. terrestris, although the impetus given to this investigation, Guido Pagliano the last species, in comparison to B. lucorum, remains for the determination of some of the bumble bees col- more abundant at lower elevations. Also taking into ac- lected in the years 1973-1993, and Marino Quaranta for count the differences between the different areas, the the determination of most of those collected in the years different sampling methods adopted in the mentioned 2001-2006; Adriana Angogna, Valerio Bertolo, Lilia studies, and the significant spatio-temporal variation Domeneghetti, Anna Saglia, and Maurizio Quirino for naturally found in bee populations (Roubik, 2001; Wil- having contributed to the Di.Va.P.R.A. collection with liams et al., 2001; Petanidou et al., 2008), we could in the specimens collected during the preparation of their any case put forward the hypothesis – to be verified thesis; Marco Anibaldi Ranco and Oscar Antonucci for through proper investigations – that the colonization of assistance with UTM WGS84 co-ordinates and figure the summit areas in being taking place by species once layout, respectively; the University of for the fi- confined to lower elevations; a similar phenomenon has nancial support of the research. been recently reported for B. terrestris and B. lapidarius in Scotland (Macdonald, 2001). 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